tyrannosaurany of a group of predatory dinosaurs that lived from the late Jurassic Period (about 150 million years ago) to the latest Cretaceous Period (about 65 million years ago), at which time they reached their greatest dominance. Most tyrannosaurs were large predators, with very large, high skulls approaching or well exceeding a full metre (more than three feet) in length. The best-known and largest member of the group is Tyrannosaurus rex, or T. rex. The “king of the tyrant lizards,” as its Latin name is usually translated, walked on powerfully developed hind limbs. If the animal had stood upright, it would have been more than 6.5 metres (21 feet) tall, but the usual posture was horizontal, with the body carried parallel to the ground and the tail held off the ground as a counterbalance. In this position a large adult, weighing 4,000 to 7,000 kg (about 9,000 to 15,000 pounds), could measure 14 metres long.

Form and function

Skull and dentition

The longest known tyrannosaur skull is 1.3 metres (more than 4 feet) long. The skull bones of large tyrannosaurs are often several centimetres thick and are strongly braced to each other, which suggests a resistance to the forces of biting, both inflicted upon and received from other tyrannosaurs. Engineering models, in fact, show that the bite force of T. rex would easily have been capable of ripping through a car roof, as portrayed in the 1993 motion picture Jurassic Park. The huge mouth contained some 60 teeth, which could protrude as far as 15 cm (6 inches). The crowns of the teeth were shed and regrown frequently (every 250 days or so, on the basis of microscopic lines visible within the teeth). Serrations of the teeth bear deep pocketlike recesses in which bacteria may have flourished to provide an infectious bite.

Tyrannosaur teeth are distinctive. The front teeth are small and U-shaped. The side teeth are large, and in adults they become even larger, fewer in number, and D-shaped in cross section rather than daggerlike as in most theropods, or flesh-eating dinosaurs. In juveniles the teeth are laterally compressed and serrated front and back, like those of other theropods. In mature individuals, however, the teeth fall neatly into three general classes: upper front teeth, upper side teeth, and lower jaw teeth. Gut contents and coprolites (fossilized feces) of tyrannosaurs, as well as remains of other dinosaurs preserved with tyrannosaurid bite marks, show that tyrannosaurs were voracious predators that could easily bite through skulls, pelvises, and limbs of other dinosaurs. Bite marks found on the bones of other tyrannosaurs, especially T. rex, have been interpreted by some scientists as evidence of cannibalism.

Other skeletal features

In contrast to the powerful jaws and legs, the forelimbs of tyrannosaurs were very small (less than the length of the shoulder blade), and in some forms the hands were reduced to only two digits. Although a mechanical reconstruction suggests that the musculature of the arms of T. rex and some other large tyrannosaurs could have lifted about 180 kg (400 pounds), the hands would not have been able to reach the mouth or grasp prey. The hind limb bones appear massive but are lightly constructed: the thickness of the bone wall is only about 20 percent of the bones’ diameter—a figure approaching that of many birds.

The age of individual dinosaurs and other vertebrates can be determined by counting the annual growth rings that are laid down in the long bones, in a manner somewhat analogous to counting tree rings. By using a series of bones from early growth stages to adulthood, the life history of an animal species can be reconstructed. Such studies have shown that T. rex effectively reached full size in less than 20 years—approximately the same period as for human beings. Of course, at a length of 6.5 metres and a mass of six tons, T. rex reached a much larger size than humans in 20 years. But its growth rate was not as high as that of some herbivorous dinosaurs such as the hadrosaurs (duck-billed dinosaurs), which reached full size in seven or eight years, or the sauropods (the largest plant-eating dinosaurs), which attained most of their gigantic size in 14 years or so. On the other hand, the growth rate of T. rex was higher than that of the African elephant, which has a similar mass yet a longer time to maturity. Some of the known specimens of T. rex did not quite reach full size; others do not seem to have survived long after achieving it. This may testify to the hard life of Mesozoic dinosaurs.

Although it was once thought that male and female tyrannosaurs could be distinguished by the shape of the tail vertebrae near the pelvis, this feature turns out not to be diagnostic. However, one subsequently discovered feature does establish sex. During the reproductive cycles of female birds, a layer of bone (medullary bone) is often deposited on the inner wall of the long bones. This process has been recognized in some fossils of tyrannosaurs (and of a few other dinosaurs), indicating that these specimens are female.

Hell Creek discoveries

Fossils of T. rex are found only in the Hell Creek Formation of Garfield county, Montana, and adjacent areas of the United States, in deposits dating from the Maastrichtian Age, the last time unit of the Cretaceous Period—although slightly earlier relatives such as Tarbosaurus and Raptorex are known from Asia. Found in the same deposits as T. rex are fossils of the ceratopsians (giant horned dinosaurs) on which they likely preyed. There is some question about whether tyrannosaurs killed their food or simply scavenged it. However, neither predatory nor scavenging behaviour need be excluded, since T. rex, like many large carnivores today, probably fed opportunistically, scavenging when it could and hunting when it had to. One argument for predation emphasizes T. rex’s vision. The eye sockets tend to be keyhole-shaped and directed forward, which has been taken as evidence for accurate depth perception, because the fields of view of the eyes would overlap. Other evidence supporting predation is the well-protected skull and formidable jaws. Wounds in the bones of its prey indicate that T. rex ate by using a “puncture and tear” stroke, planting its feet and using the powerful muscles of the neck and legs to anchor itself and pull flesh off bones.

Before 1980 all knowledge of T. rex was based on only four skeletons, none very complete. The Latin name was given to the first specimen by American paleontologist Henry Fairfield Osborn in 1905 and was based on partial specimens collected from the Hell Creek Formation by renowned fossil hunter Barnum Brown. Remains found by Brown are on display at the Carnegie Museum of Natural History in Pittsburgh, Pa., the American Museum of Natural History in New York City, and the Natural History Museum in London. Since 1980 more than two dozen other specimens of T. rex have been discovered in western North America, some very complete; however, some are in private collections and are therefore lost to science and education. Two of the best specimens, consisting of almost complete adult skeletons, were unearthed in 1990. One, the 85-percent-complete “Wankel” T. rex, is on display at the Museum of the Rockies in Bozeman, Mont., and the other, the 90-percent-complete “Sue,” is displayed at the Field Museum in Chicago. Other T. rex specimens are mounted at other natural history museums in North America, such as the Denver Natural History Museum, the University of California Museum of Paleontology in Berkeley, the Natural History Museum of Los Angeles County, and the Royal Tyrrell Museum in Drumheller, Alta., near Dinosaur Provincial Park.

In 2000 five T. rex specimens were discovered in the Hell Creek Formation. Several are now on display at the Museum of the Rockies; one of them, the “B-rex,” preserves soft tissues and also medullary bone that indicates the specimen was female. The soft tissues preserve transparent, flexible, hollow blood vessels that contain small round microstructures highly reminiscent in structure of red blood cells. The preservation of these structures is one of the most amazing features of the entire known fossil record.

Classification

Tyrannosaurs are generally divided into the large but more lightly built and slightly earlier albertosaurines and the still larger, more robust, and later tyrannosaurines. Most tyrannosaurs are known from the latest Cretaceous, but some basal forms are now known from the Early Cretaceous and even the Late Jurassic, though these earlier forms share few features with their later, well-known relatives. Guanlong, an animal about three metres long from the Late Jurassic of Xinjiang province, western China, is the earliest well-known member of the group; it has some primitive and unique features—the most notable being a complex skull crest consisting of a hollow bone running atop the midline of its skull. Dilong, an early tyrannosaur 1.5 metres (5 feet) long from the spectacular Liaoning deposits of northeastern China, is preserved with a covering of simple, filamentous “protofeathers” like those seen on many other Early Cretaceous theropod dinosaurs. Eotyrannus, from Early Cretaceous deposits on Britain’s Isle of Wight, is also lightly built and relatively small (some 4.5 metres, or 15 feet, long). These three tyrannosaurs are so primitive that they retain three fingers on their hands.

Several small tyrannosaur fossils from the latest Cretaceous of western North America were once assigned to separate taxa, but most scholars now consider them to be merely young tyrannosaurs. For example, specimens once given the names Nanotyrannus and Stygivenator are now considered to be juvenile tyrannosaurs, and the former Dinotyrannus is now seen as a subadult T. rex. T. rex is the only tyrannosaur known from the late Maastrichtian Age (i.e., the latest Cretaceous Period) in North America. As is mentioned above, Tarbosaurus is a slightly earlier and very similar form from the latest Cretaceous of Mongolia.

Formally described in 2009, a new genus of earlier and smaller tyrannosaur, Raptorex kriegsteini, is based on a single specimen from the Early Cretaceous of the Inner Mongolia Autonomous Region. It stood roughly 3 metres (10 feet) tall, and its weight was roughly 40 to 70 kg (90 to 150 pounds), about one-hundredth the weight of Tyrannosaurus. Although R. kriegsteini is small for a tyrannosaur, as most of the basal ones are, its body plan—with smaller, D-shaped tooth cross sections and short arms with reduced fingers—closely resembles a reduced version of T. rex.

Tyrannosaurs were long thought to be one of the carnosaurs (“flesh-eating lizards”), related to other large theropods such as the allosaurs. These resemblances have proved to be superficial, related to large size alone. Today tyrannosaurs are considered to be gigantic members of the coelurosaurs (“hollow-tailed lizards”), a group largely composed of smaller, more-gracile forms. Frequently they have been related to the largely toothless, ostrichlike ornithomimids, mainly because tyrannosaurs and ornithomimids share a peculiar foot with “pinched” foot bones. Tyrannosaurs may turn out to be closely related to the dromaeosaurs, the “raptors” of Jurassic Park, though evidence for this hypothesis is as elusive as any other.